Zootaxa 1553: 47–58 (2007) www.mapress.com / zootaxa/

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ZOOTAXA

The Ixodes (Acari: Ixodidae) of Mexico: parasite-host and host-parasite checklists CARMEN GUZMÁN-CORNEJO1, RICHARD G. ROBBINS2 & TILA M. PÉREZ1 1

Colección Nacional de Ácaros, Instituto de Biología, Universidad Nacional Autónoma de México, 3er Circuito Exterior s/n, Ciudad Universitaria, Copilco, Coyoacán, México D.F., C.P. 04510. E-mail:[email protected] 2 DPMIAC/AFPMB, Walter Reed Army Medical Center, Washington, DC 20307-5001.

Abstract Parasite-host and host-parasite checklists are provided for all species of Ixodes known from Mexico; host and locality data are from specimens housed in the Colección Nacional de Ácaros, Instituto de Biología, Universidad Nacional Autónoma de México, and from literature. Six Ixodes species (I. brunneus, I. conepati, I. dentatus, I. eadsi, I. guatemalensis, I. texanus) are newly recorded from Mexico; in addition, 17 new locality records are presented for eight species (I. affinis, I. boliviensis, I. luciae, I. rubidus, I. scapularis, I. spinipalpis, I. tancitarius, I. woodi), and eight new host records are given for five species (I. affinis, I. boliviensis, I. rubidus, I. spinipalpis, I. tancitarius). Key words: Ixodidae, Ixodes, Mexico, checklists, hosts, distribution.

Introduction Ixodes Latreille is the largest genus of ticks in the world, currently comprising 243 species (Guglielmone et al. 2006), more than one quarter of the global tick fauna. In all life history stages, members of this genus possess an anal groove that curves anterior to the anus, forming an arch. Several Ixodes species are of major medical and veterinary importance, having been implicated in the transmission of zoonotic disease agents (Sonenshine et al. 2002; Goodman et al. 2005). Despite the predominance of this genus in the Northern Hemisphere, relatively few species of Ixodes have been reported from Mexico. Hoffmann and López-Campos (2000) recorded 18 Ixodes species associated with Mexican birds and mammals, while Whitaker and Morales-Malacara (2005), working only with mammals, reported 14 species. As part of a project to catalog the tick species deposited in the Colección Nacional de Ácaros (CNAC), Instituto de Biología, Universidad Nacional Autónoma de México (IBUNAM), we analyzed all specimens belonging to the genus Ixodes. Initially, the CNAC chiefly consisted of Dr. Anita Hoffmann’s personal collection, which was donated to IBUNAM in 1992, and in 1997 was officially designated the National Collection, housing a great variety of mites and ticks from Mexico. The objective of this paper is to compile parasite-host and host-parasite checklists for all known Mexican species of Ixodes and to report new host, distribution and taxonomic data derived from specimens deposited in the CNAC.

Accepted by H. Klompen: 12 Jun. 2007; published: 20 Aug. 2007

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Materials and methods To construct our checklists, we first conducted a number of online bibliographic searches, using such databases as AGRICOLA, BIOSIS, CAB Abstracts, Medline, and the Zoological Record; these were supplemented by information contained in the CNAC database (Biota version 1.6.1). Unidentified Ixodes specimens were determined using conventional morphological keys to this genus (Cooley & Kohls 1942; Cooley 1943; Cooley & Kohls 1945; Kohls 1956; Kohls & Clifford 1966; Keirans & Clifford 1978; Robbins & Keirans 1992; Durden & Keirans 1996), and the resulting records were entered into the CNAC database. The parasite-host checklist comprises previously published records obtained through bibliographic searches and, when available, new records resulting from examination of CNAC specimens. Tick collection records are presented in the following order: state (capitalized and in boldface), number and sex or stage of tick specimens, locality, date, host species name (updated, if necessary), and, for published records, references (in brackets) and depository (in parentheses). Some new records are accompanied by geographic coordinates, and all have been assigned CNAC accession numbers (in parentheses). Abbreviations: NA, no information available; ENV, École Nationale Vétérinaire, Toulouse, France; and RML, Rocky Mountain Laboratories. All RML specimens are now in the United States National Tick Collection (USNTC), Institute of Arthropodology and Parasitology, Georgia Southern University, Statesboro, USA. The host-parasite checklist presents host species alphabetically within each vertebrate order and family. Host names have been updated to accord with those of Ramírez-Pulido et al. (1996) and Ceballos and Oliva (2005) for mammals, and the American Ornithologists’ Union (1998) for birds.

Results Our checklists contain host and distributional data for 26 Ixodes species parasitizing 28 species of vertebrates (five birds and 23 mammals) in Mexico. Eleven earlier records were corroborated from specimens in CNAC; the remainder are from literature. The species I. brunneus, I. conepati, I. dentatus, I. eadsi, I. guatemalensis and I. texanus represent new records for Mexico. In addition, we provide 17 new locality records for eight tick species (I. affinis, I. boliviensis, I. luciae, I. rubidus, I. scapularis, I. spinipalpis, I. tancitarius and I. woodi), and eight new host records for five species (I. affinis, I. boliviensis, I. rubidus, I. spinipalpis, and I. tancitarius). The records for I. granulatus are erroneous.

Parasite-host list Ixodes affinis Neumann Previous records CHIAPAS: 1&, Ocozocoautla, XII-1946, Nasua narica [Kohls & Rogers 1953]; same data except 1%, 7&, Mazama americana; NA, Mapastepec, “deer” [Hoffmann 1962]. New records CHIAPAS: 14&, 6%, NA, NA, (CNAC002247); 3&, 1%, Mapastepec, 29-II-1954, “deer” (CNAC002159). YUCATÁN: 1&, NA, Bos taurus (recorded as “bovine”), (CNAC002293). Comments: The earlier record from Mapastepec, cited by Hoffmann (1962), lacks a date, and the collector was recorded as Macías. Our new record includes a date, and the collector was a Dr. Tort.

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Ixodes angustus Neumann Previous records CHIAPAS: NA, Cerro Tzontehuitz, 13 km NE San Cristóbal de las Casas, San Juan Chamula, 24-X1993, Reithrodontomys microdon microdon [Estébanes-González & Cervantes 2005]; NA, Cerro Tzontehuitz, 13 km NE San Cristóbal de las Casas, San Juan Chamula, 24-X-1993, Microtus guatemalensis [EstébanesGonzález & Cervantes 2005]. COAHUILA: 10&, Sabinas, I-1944, nest of Neotoma sp. [Hoffmann 1962] (RML21723). OAXACA: NA, 3 km SE Totontepec, Totontepec, 5-XI-1990, Peromyscus melanocarpus [Estébanes-González & Cervantes 2005]; NA, 11 km SW Esperanza to San Isidro, Santiago Comaltepec, 9IX-1990, Sorex sp. [Estébanes-González & Cervantes 2005]; NA, 1 km N Esperanza, Santiago Comaltepec, Distrito Ixtlán, 8-XII-1989, Oryzomys alfaroi [Estébanes-González & Cervantes 2005]; NA, San Martín Caballero, Distrito Teotitlán, San José Tenango, 24-V-1994, “mouse” [Estébanes-González & Cervantes 2005]. Comments: We examined the specimens cited by Estébanes-González & Cervantes (2005) from Reithrodontomys microdon microdon and Peromyscus melanocarpus and have concluded that they do not correspond to I. angustus, thus casting doubt on all determinations of this species by these authors. We have not seen the specimens studied by Hoffmann (1962), but in light of this species’ largely North Temperate distribution (Robbins & Keirans 1992), we can only tentatively include it in the Mexican tick fauna. Ixodes bequaerti Cooley and Kohls Previous record CHIAPAS: 1&, Catharinus (sic), 4-V-1942, NA [Cooley & Kohls, 1945]. Ixodes boliviensis Neumann Previous records CHIAPAS: 1&, Planta de Luz, Huixtla, VI-1945, Canis familiaris (recorded as “dog”) [Hoffmann 1962] (CNAC002165); 2&, Santa Rosa, Comitán, 14-VI-1937, NA [Hoffmann 1962] (CNAC002162). GUERRERO: 3&, Atoyac, Panthera onca, Homo sapiens [Neumann 1906] (ENV). NAYARIT: 1&, NA, “deer” [Cooley & Kohls 1945] (RML19820). OAXACA: 1&, Teotila, Cuicatlán, Oaxaca, II-1953, NA [Hoffmann 1962] (CNAC002160); 3&, Yaviche, VII-1950, Canis familiaris (recorded as “dog”) [Hoffmann 1962] (CNAC002164). QUINTANA ROO: 1&, Bacalar, 29-VIII-1939, “fox” [Hoffmann 1962] (CNAC002163). Comments: The specimens from Guerrero and Nayarit were originally reported as Ixodes bicornis Neumann, which was relegated to a junior synonym of I. boliviensis by Kohls (1956). New records CHIAPAS: 11&, Finca Prusia, 12-XII-1944, NA (CNAC005123). OAXACA: 1&, Tarabundí (17° 36' 34”N, 96° 18' 55”W), 6-XII-1948, “coati” (CNAC002252); 2&, Tarabundí, 10-XII-1948, “pheasant” (CNAC002253). VERACRUZ: 1&, 0.8 km NE Las Minas (19° 41' 27”N, 97° 8' 47”W), 10-V-1963, Sciurus sp. (CNAC002352); 1&, Rancho la Encantada, San Andrés Tuxtla, 5-XI-2002, NA (vegetation) (CNAC005059); 1&, Cerro el Vigía, Santiago Tuxtla, 7-IV-1967, Homo sapiens (recorded as “man”) (CNAC005075). Ixodes brunneus Koch New record MEXICO D. F.: 1&, Chapultepec, 4-XI-1950, NA (CNAC005161). Ixodes conepati Cooley and Kohls New records COAHUILA: 1&, Cueva la Manga, Rancho San Judas Tadeo, Acuña (29° 21' 47”N; 101° 1' 47”W), 2-

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IV-2005, NA (cave) (CNAC002353); 1&, Cueva Popo de Oso, Zaragoza (28° 38' 4”N; 101° 7' 17”W), 24-XI2006, NA (cave) (CNAC002362). Ixodes cookei Packard Previous records NUEVO LEÓN: 15 NN, 5 LL, 8.0 km N, 4.1 km W San Josecito, 22-II-1990, Bassariscus astutus [Montiel-Parra et al. 2007] (CNAC); 2 LL, 10.8 km S, 10.8 km E San Josecito, 12-VI-1991, Peromyscus sp. [Montiel-Parra et al. 2007] (CNAC). VERACRUZ: 1%, Área Natural Protegida San Juan del Monte, 3.2 km SW, Las Vigas de Ramírez, 25-VIII-2003, Urocyon cinereoargenteus [Montiel-Parra et al. 2007] (CNAC). Ixodes cuernavacensis Kohls and Clifford Previous record MORELOS: 1&, Cuernavaca, 30-V-1961, Streptoprocne semicollaris [Kohls & Clifford 1966] (RML 37920). Ixodes dampfi (Cooley) Previous records ESTADO DE MÉXICO: 5&, Valle de México, Chalco, IV-1932, Geomys sp. [Cooley 1943] (RML 20165); Collection of Dr. A. Dampf (Mexico City); National Museum of Natural History (formerly United States National Museum), Washington, DC; and Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA. Ixodes dentatus Marx New record NUEVO LEON: 2&, 2%, El Potosí (24° 49' 42”N; 100° 20' 26”W), 22-V-1963, Sylvilagus sp. (CNAC002366). Ixodes eadsi Kohls and Clifford New record PUEBLA: 1&, 6 km E Totimehuacán, 13-III-1965, Liomys sp. (CNAC002287). Ixodes granulatus Supino Previous records OAXACA: NA, 1 km S, 0.5 km E Vista Hermosa, Santiago Comaltepec, 20-X-1989, Peromyscus mexicanus totontepecus [Estébanes-González & Cervantes 2005]; NA, 11 km SW La Esperanza to San Isidro, Santiago Comaltepec, 30-V-1990, Oryzomys chapmani chapmani [Estébanes-González & Cervantes 2005]. Comments: Ixodes granulatus is an exclusively Asian species, ranging from Japan through Southeast Asia and westward to India (Petney & Keirans 1994). The larvae from Peromyscus mexicanus totontepecus do not correspond to this species, and other purported Mexican “I. granulatus” should also be redetermined. Ixodes guatemalensis Kohls New records GUERRERO: 6&, Omiltemi (17° 33' 24”N; 99° 41' 8”W), 4-IV-1963, Sciurus sp. (CNAC002325); 2&, Omiltemi, 5-IV-1963, Sciurus sp. (CNAC002359); 2&, Omiltemi, IV-1963, Sciurus sp. (CNAC002356); 1&, Omiltemi, 2-IV-1963, Sciurus sp. (CNAC002360); 3&, 1.6 km NW Omiltemi, 13-IV-1963, Sciurus sp. (CNAC002335); 18&, 1.6 km NW Omiltemi, 15-IV-1963, Sciurus sp. (CNAC002367). VERACRUZ: 1&, 0.8 km NE Las Minas, 5-V-1963, Sciurus sp. (CNAC002337).

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Ixodes loricatus Neumann Previous record TABASCO: NA, Frontera, Ateles geoffroyi [Nuttall & Warburton 1911]. Comments: The locality cited by Nuttall and Warburton (1911), “Tabasco de la Frontera,” is probably the city of Frontera, in Tabasco. Ateles “melanochoerus” (= melanocercus or melanochir) is a junior synonym of A. geoffroyi. Ixodes luciae Sénevet Previous records CHIAPAS: 1&, 1%, Finca Germania, 27-XII-1943, Didelphis sp. [Vázquez 1946] (CNAC000569, CNAC000570). TABASCO: 3&, Frontera, V, “large opossum” [Nuttall 1910; Nuttall & Warburton 1911]. Comments: The specimens from Finca Germania were originally reported as Ixodes scuticrenatus Vázquez, while those from Frontera were reported as Ixodes loricatus variety spinosus Nuttall; both these names are junior synonyms of I. luciae (Camicas et al. 1998). The locality cited by Nuttall and Warburton (1911), “Tabasco de la Frontera,” is probably the city of Frontera, in Tabasco. New records COLIMA: 1&, 1%, La Barragana, 12-IX-1978, Didelphis marsupialis (CNAC002316). VERACRUZ: 4%, 0.8 km NE Las Minas, 9-V-1963, Didelphis sp., (CNAC002354). Comments: The specimens from Colima were previously identified by R. H. Manzanilla L.; however, this is a new locality record for Mexico. Ixodes mexicanus Cooley and Kohls Previous records MICHOACÁN: 1&, Cerro Tancítaro, VIII-1940, Campylorhynchus gularis [Cooley & Kohls 1942]; same data except 17-VII-1940, Junco phaeonotus (RML 17469). Ixodes murreleti Cooley and Kohls Previous record BAJA CALIFORNIA SUR: 1&, Isla los Coronados, 4-IV-1940, “Scripps’ [= Xantus’s] murrelet,” Synthliboramphus hypoleucus (formerly Endomychura hypoleucus) [Cooley & Kohls 1945] (RML 17799). Ixodes pacificus Cooley and Kohls Previous record BAJA CALIFORNIA: NA, NA [Bishopp & Trembley 1945]. Comments: Bishopp and Trembley (1945) stated that Ixodes ricinus californicus Banks occurs along the Pacific coast of North America from Mexico to British Columbia; however, this name is a junior synonym of I. pacificus (Cooley & Kohls 1945). Hoffmann (1962), citing Bishopp and Trembley (1945) but using the nomenclature of Cooley and Kohls (1945), referred Mexican records of I. pacificus to Baja California. Ixodes rubidus (Neumann) Previous records GUANAJUATO: 1&, 3NN, NA, Bassariscus astutus [Neumann 1911; Nuttall & Warburton 1911; Cooley & Kohls 1945] (ENV). New record GUERRERO: 3&, 36NN, Omiltemi, 8-IV-1963, Urocyon sp. (CNAC002364). Comments: The host of these specimens must be Urocyon cinereoargenteus because only one species of this genus occurs in Mexico (Ceballos & Oliva, 2005).

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Ixodes scapularis Say Previous records JALISCO: NA, Purificación, “cattle” [Chavarría, 1941]. OAXACA: NA, Finca “El Sinaí,” Santos Reyes Nopala, Distrito Juquila (16° 06´N, 97° 09´W), NA [Marín & Hoffmann 2002]. TAMAULIPAS: 1&, 1%, west of Tampico, NA [Hooker et al. 1912]; same data except Canis familiaris, “cattle” [Macías Valadez, 1923]. Comments: Chavarría (1941) used the name Ixodes ricinus variety scapularis Say, which is a junior synonym of I. scapularis (Camicas et al. 1998). Hoffmann (1962), citing Macías Valadez (1923), stated that Felis pardalis, now Leopardus pardalis, is a host of I. scapularis; however, this record is from Costa Rica. New record COAHUILA: 9&, 1%, Las Herminias, Zaragoza, 23-II-1975, Bos taurus (recorded as “bovine”) (CNAC002166). Comments: These specimens were previously identified by A. de la Torre; however, this is a new locality record for Mexico. Ixodes sinaloa Kohls and Clifford Previous records NAYARIT: 1&, 32 km SE Tepic, 18-VII-1963, Oryzomys couesi mexicanus [Kohls & Clifford 1966] (RML 46302). SINALOA: 1&, 5 km SW San Ignacio, 20-VII-1963, Liomys pictus escuinapae [Kohls & Clifford 1966] (RML 39478); same data except 2&, 19-VII-1965 (RML 39475, RML 39477); 1&, 1 km NE Santa Lucia, 24-VII-1965, Liomys pictus escuinapae [Kohls & Clifford 1966] (RML 39479); 1&, 4.8 km NE Copala, 12-VII-1963, Liomys pictus escuinapae [Kohls & Clifford 1966] (RML 46296); 1&, 6.4 km SW Copala, 2-VIII-1960, Liomys pictus escuinapae [Kohls & Clifford 1966] (RML 46294). Ixodes spinipalpis Hadwen and Nuttall Previous record MORELOS: 1&, km 43 Carretera México-Cuernavaca, 26-VI-1949, Romerolagus diazi [Hoffmann 1962] (CNAC005180). Comments: Hoffmann’s (1962) record was for Ixodes neotomae Cooley; however, Norris et al. (1997) argued that this taxon is conspecific with, and therefore a junior synonym of, I. spinipalpis. New records GUERRERO: 1N, Omiltemi, (17° 33' 24”N; 99° 41' 8”W), 2-IV-1963, NA (CNAC002358); 5NN, Omiltemi, 3-IV-1960, Peromyscus sp. (CNAC002331); 6NN, Omiltemi, 3-IV-1963, Peromyscus sp. (CNAC002332); 7NN, Omiltemi, 6-IV-1963, Peromyscus sp. (CNAC002333); 3NN, Omiltemi, 5-IV-1963, Sciurus sp. (CNAC002341); 2NN, 1.6 km NW Omiltemi, 2-IV-1963, Peromyscus sp. (CNAC002355). MORELOS: 2NN, Zempoala, Peromyscus sp. (CNAC002326). VERACRUZ: 1N, 0.8 km NE Las Minas, 2V-1963, Peromyscus sp. (CNAC002350). Ixodes tamaulipas Kohls and Clifford Previous record TAMAULIPAS: 1&, Rancho del Cielo southeast of Tamaulipas State, 8-VIII-1963, Sciurus deppei negligens [Kohls & Clifford 1966] (RML 39326). Ixodes tancitarius Cooley and Kohls Previous record MICHOACÁN: 1&, Cerro Tancítaro, 5-VII-1941, Reithrodontomys sp. [Cooley & Kohls 1942] (RML 19236).

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New record VERACRUZ: 1&, 0.8 km NE Las Minas, 2-V-1963, Peromyscus sp. (CNAC002336). Ixodes texanus Banks New records GUERRERO: 1&, 1N, Omiltemi (17° 33' 24”N; 99° 41' 8”W), 6-IV-1963, Bassariscus sp. (CNAC002370). Ixodes tovari Cooley Previous records GUANAJUATO: 2&, 2%, Torrecilla and Comontoso, “hares” [Cooley 1945] (RML 21620–21621); NUEVO LEON: 19 “adults,” 9%, 7NN, 2LL, Bravo, “hares” [Cooley 1945] (RML 21618, 21623–21625, 21628). Ixodes woodi Bishop Previous records COAHUILA: 3&, 1N, Sabinas (27o 50’N; 101o 9’W), I-1944, Neotoma micropus [Robbins & Keirans 1987] (RML 47170). TAMAULIPAS: Rancho del Cielo, NA [Robbins & Keirans 1987] (RML correspondence files). New record MORELOS: 1&, Zempoala (19° 3' 11”N; 99° 18' 44”W), 23-III-1963, Neotoma sp. (CNAC002340)

Host-parasite list AVES APODIFORMES Apodidae Streptoprocne semicollaris (De Saussure)

Ixodes cuernavacensis

CHARADRIIFORMES Alcidae Synthliboramphus hypoleucus (Xantus de Vesey)

Ixodes murreleti

GALLIFORMES “pheasant”

Ixodes boliviensis

PASSERIFORMES Emberizidae Junco phaeonotus Wagler Troglodytidae Campylorhynchus gularis Sclater

Ixodes mexicanus Ixodes mexicanus

MAMMALIA ARTIODACTYLA

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Bovidae Bos taurus L. “cattle” Cervidae “deer” Mazama americana (Erxleben) CARNIVORA Canidae “fox” Canis familiaris L. Urocyon cinereoargenteus (Schreber) Urocyon sp. Felidae Panthera onca (L.) Procyonidae Bassariscus astutus (Lichtenstein)

Ixodes affinis Ixodes scapularis Ixodes scapularis I. affinis I. boliviensis Ixodes affinis

Ixodes boliviensis Ixodes boliviensis Ixodes scapularis Ixodes cookei Ixodes rubidus Ixodes boliviensis

Bassariscus sp. “coati” Nasua narica (L.)

Ixodes cookei Ixodes rubidus Ixodes texanus Ixodes boliviensis Ixodes affinis

DIDELPHIMORPHIA Didelphidae Didelphis marsupialis L. Didelphis sp. “opossum”

Ixodes luciae Ixodes luciae Ixodes luciae

LAGOMORPHA Leporidae Lepus sp. (“hares”) Romerolagus diazi (Ferrari-Pérez) Sylvilagus sp.

Ixodes tovari Ixodes spinipalpis Ixodes dentatus

PRIMATES Atelidae Ateles geoffroyi Kuhl Hominidae Homo sapiens L. RODENTIA “mouse”

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Ixodes loricatus Ixodes boliviensis

Ixodes sp.

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Geomyidae Geomys sp. Heteromyidae Liomys pictus escuinapae J. A. Allen Liomys sp. Muridae Microtus guatemalensis Merriam Neotoma micropus Baird Neotoma sp. Oryzomys alfaroi (J. A. Allen) Oryzomys couesi (Alston) Peromyscus melanocarpus Osgood Peromyscus sp.

Reithrodontomys microdon microdon Merriam Reithrodontomys sp. Sciuridae Sciurus deppei negligens Nelson Sciurus sp.

SORICOMORPHA Soricidae Sorex sp.

Ixodes dampfi Ixodes sinaloa Ixodes eadsi Ixodes sp. Ixodes woodi Ixodes angustus Ixodes woodi Ixodes sp. Ixodes sinaloa Ixodes sp. Ixodes cookei Ixodes spinipalpis Ixodes tancitarius Ixodes sp. Ixodes tancitarius Ixodes tamaulipas Ixodes boliviensis Ixodes guatemalensis Ixodes spinipalpis

Ixodes sp.

Discussion Mexico is considered a megadiverse country, so it is not surprising that its invertebrate fauna has been incompletely cataloged. Taxonomic knowledge of Mexican ticks remains limited, particularly in the case of Ixodes, which is currently represented by just 26 species, i.e., only 10.7% of the known species in this genus (Mexican records of I. granulatus are erroneous). By contrast, 35 species of Ixodes are known from the United States and Canada, despite the temperate and boreal conditions that prevail over most of northern North America. Furthermore, in the United States, species of Ixodes are known to parasitize reptiles, birds and mammals (Cooley & Kohls 1945, Durden & Keirans 1996, Wright et al. 1998, Tälleklint-Eisen & Eisen 1999, and Eisen et al. 2001), whereas in Mexico only birds and mammals have been reported as hosts, and among mammals, rodents predominate, hosting 11 Ixodes species. Of some 1,050 species of birds and 452 species of mammals found in Mexico, only 0.47% and 5.1%, respectively, have been recorded as hosts for Ixodes ticks. Clearly, there is a pressing need for extensive parasitological sampling of Mexico’s approximately 2,306 species of terrestrial vertebrates (Flores-Villela 1993, Ramírez-Pulido et al. 1996, Ceballos & Márquez 2000, Flores-Villela & Canseco-Márquez 2004, Ceballos & Oliva 2005). Currently, Bassariscus astutus, Bos taurus and Canis familiaris are each known to host two Ixodes species; the rodent genera Peromyscus and Sciurus both host three Ixodes species, but in these cases more than one host species may be involved. To date, research on Mexican Ixodes has largely been conducted outside Mexico (Cooley & Kohls 1942, Kohls & Clifford 1966), with the result that most literature records are based on specimens in foreign collecTHE IXODES (ACARI: IXODIDAE) OF MEXICO

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tions, such as the RML and ENV. Currently, the CNAC contains representatives of only 15 of the 26 Ixodes species known to occur in Mexico. We have corroborated the identity of three of the CNAC species (I. boliviensis, I. luciae, and I. spinipalpis); 12 more, including six new national records, resulted from the present study. All determinations were based on adults except for I. spinipalpis, which was based on nymphs. The first report of Ixodes cookei in Mexico was published separately (Montiel-Parra et al. 2007). Species of Ixodes have been collected in 21 of the 32 states of Mexico; the highest number of species has been recorded in Veracruz (six species), followed by Chiapas and Guerrero (five species), and Coahuila (four species). The most widespread tick species in Mexico appear to be I. boliviensis (Chiapas, Guerrero, Nayarit, Oaxaca, Quintana Roo and Veracruz), I. luciae (Chiapas, Colima, Tabasco, Veracruz), and I. scapularis (Coahuila, Jalisco, Oaxaca, Tamaulipas). The broad range of I. boliviensis also corresponds with the highest number of associated hosts. Additional systematic research on Mexican ticks is urgently needed, not only to catalog this group, but also to augment regional studies of ecology, biogeography, and host-parasite coevolution. Moreover, taxonomic elucidation of the Mexican tick fauna will substantively contribute to investigations of zoonotic infections, such as Lyme disease, that merit increased attention in Mexico.

Acknowledgments We warmly thank the Dirección General de Asuntos del Personal Académico (DGAPA), Universidad Nacional Autónoma de México, for the award of a postdoctoral scholarship to the senior author. We also thank Peter Sprouse for donation of specimens of Ixodes conepati to CNAC.

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